Ecological memory preserves phage resistance mechanisms in bacteria

Defense mechanisms against pathogens are prevalent in nature, and their maintenance is critical for long-term survival of a species. Such mechanisms, which include CRISPR-mediated immunity in bacteria and the R genes in plants, carry substantial costs to organisms and can be rapidly lost when pathogens are eliminated. How a species preserves its molecular defenses despite their costs, in the face of variable pathogen levels, and across an ecology of localized patches remains a major unsolved problem in epidemiology and evolutionary biology. Using techniques of game theory and non-linear dynamical systems, we show that by maintaining a non-zero failure rate of immunity, hosts sustain sufficient levels of pathogen across an ecology to select against loss of the defense. This resistance switching strategy is evolutionarily stable, and provides a powerful evolutionary mechanism that maintains host-pathogen interactions and enables co-evolutionary dynamics in a wide range of systems.